Paleobiología de los artrópodos del ámbar cretácico de El Soplao (Cantabria, España) = Arthropod paleobiology of the cretaceous amber from El Soplao (Cantabria, Spain)

[spa] El yacimiento de ámbar de El Soplao (Rábago, Cantabria), de edad Cretácico Inferior (hace unos 110 millones de años), ha proporcionado a día de hoy más de 500 bioinclusiones de artrópodos. El estudio morfológico y sistemático de los artrópodos del ámbar de El Soplao se ha centrado en rafidiópteros, neurópteros y algunos grupos de dípteros, himenópteros y arañas, y ha permitido realizar inferencias paleoecológicas, paleobiogeográficas y paleoetológicas. Estos datos han complementado los datos geológicos y otros datos paleontológicos (como los paleobotánicos), que, en suma, han conducido a una reconstrucción paleoambiental. La presente Tesis Doctoral se basa en un total de once artículos. Las plantas productoras de resina que dieron origen al ámbar de El Soplao, coníferas pertenecientes al menos a la familia extinta Cheirolepidiaceae y a otro grupo diferente por determinar, se desarrollarían cerca de la costa marina. Los incendios, como parte integrante de la dinámica del ecosistema, tuvieron un papel determinante en la producción y posterior acumulación de resina que dio lugar al yacimiento, provocando la secreción traumática de resina (en la que los insectos barrenadores de madera pudieron haber colaborado) y eliminando la cobertura vegetal del suelo del bosque. Ello facilitó que el agua de escorrentía erosionara el suelo y movilizara la resina depositada primariamente. Esta resina sufrió un corto transporte y se depositó de un modo paraauctóctono en las áreas de enterramiento definitivo, al no existir indicios de reelaboración ni retrabajamiento del ámbar. Estas áreas donde la resina quedó enterrada y fosilizó corresponden a ambientes de marjal costero o de estuario asociado a una llanura deltaica. Algunas masas de resina presentaron un largo periodo de exposición en las aguas salobres o marinas antes de enterrarse. Pese a hallarse en una fase de estudio inicial, la diversidad identificada de artrópodos en el ámbar de El Soplao es elevada, y en algunas casos ha brindado el registro más antiguo conocido de ciertos taxones. En un total de 549 inclusiones de artrópodos catalogadas, se han determinado 13 órdenes de hexápodos repartidos en unas 40 familias, cuatro órdenes de arácnidos y uno de crustáceos. La paleofauna de artrópodos de El Soplao muestra una serie de singularidades, tanto cuantitativas como cualitativas, en comparación con el resto de paleofaunas conocidas de los ámbares cretácicos. Estas singularidades paleofaunísticas tanto a nivel cuantitativo como cualitativo en el ámbar de El Soplao, aparte de los factores tafonómicos y teniendo además en cuenta los hallazgos anteriores en el ámbar de Peñacerrada I, podrían reflejar un cierto carácter endémico de los ecosistemas ibéricos durante el Albiense debido a la insularidad que la Placa Ibérica adquirió en el Jurásico Inferior. Existen tres especies de insectos comunes a los ámbares de El Soplao y Peñacerrada I (= Moraza). Ello sugiere una edad muy similar para ambos yacimientos de la Cuenca Vasco‐Cantábrica. Además, se han advertido ciertas similitudes entre las paleofaunas del ámbar de España (El Soplao y Peñacerrada I) (Albiense) y las del ámbar del Líbano (Barremiense‐ Aptiense), aunque se hallen espaciadas en el tiempo por varios millones de años. No obstante, es necesario aumentar las colecciones de los ámbares españoles para dar mayor fundamento a estas observaciones, así como revisar las dataciones del ámbar del Líbano. La elevada proporción de sininclusiones y muestras que brindan información paleoetológica o acerca de interacciones paleobióticas, tanto intraespecíficas como interespecíficas, dan al ámbar de El Soplao un elevado valor paleoecológico. Existen casos de posible enjambramiento, de cópula, de camuflage/defensa, de relación planta‐insecto, de parasitismo, de posible hematofagia y de posible relación depredador‐presa.[eng] The El Soplao amber outcrop (Rábago, Cantabria, Northern Spain), Early Cretaceous in age (about 110 Mya), has currently provided more than 500 arthropod inclusions. This PhD dissertation aimed to achieve in‐depth knowledge of the paleobiology associated with the arthropods from the El Soplao amber outcrop, allowing for a preliminary comparison with primary Cretaceous amber outcrops. This was possible only after a thorough systematic treatment of El Soplao arthropod inclusions that inferred their paleoautoecological, paleobiogeographical and paleoethological contexts to, ultimately, glean a holistic understanding of the paleoenvironmental setting. The resin‐producing plants responsible for El Soplao amber are conifers belonging to the extinct family Cheirolepidiaceae and, at least, another unknown conifer group. These trees would have lived close to the paleo‐coastline. Paleofires played a leading role in the production and accumulation of resin. The areas where the resin was buried and fossilized correspond to marshy or delta‐estuarine environments. Some resin pieces had a long exposure time to marine or brackish waters before being buried. Identified arthropod diversity is high, and the outcrop has provided the oldest known records for certain taxa. Thirteen hexapod orders (composed by about 40 families), four arachnid orders, and one crustacean order have been determined. The arthropod fauna from El Soplao is quantitatively and qualitatively singular when compared to faunas from other known Cretaceous ambers. Three insect species occur in both the El Soplao and Peñacerrada I (=Moraza) outcrops. The shared species between these two Basque‐Cantabrian Basin outcrops suggest they are similar in age. Moreover, similarities have been noted between the paleofauna from the Spanish amber outcrops (El Soplao and Peñacerrada I) (Albian) and the paleofauna from Lebanon (Barremian‐Aptian), despite these outcrops are separated in time by several million years. El Soplao amber is valuable in the abundance of syninclusions and samples that provide information about paleoethology and paleobiotic interactions. There are instances of possible swarming, mating, camouflage/defense, plant‐insect relationship, parasitism, possible hematophagy and possible predator‐prey relationship. An expanded English summary is provided at the end of the dissertation

Paleobiología de los artrópodos del ámbar cretácico de El Soplao (Cantabria, España) = Arthropod paleobiology of the cretaceous amber from El Soplao (Cantabria, Spain)

El yacimiento de ámbar de El Soplao (Rábago, Cantabria), de edad Cretácico Inferior (hace unos 110 millones de años), ha proporcionado a día de hoy más de 500 bioinclusiones de artrópodos. El estudio morfológico y sistemático de los artrópodos del ámbar de El Soplao se ha centrado en rafidiópteros, neurópteros y algunos grupos de dípteros, himenópteros y arañas, y ha permitido realizar inferencias paleoecológicas, paleobiogeográficas y paleoetológicas. Estos datos han complementado los datos geológicos y otros datos paleontológicos (como los paleobotánicos), que, en suma, han conducido a una reconstrucción paleoambiental. La presente Tesis Doctoral se basa en un total de once artículos. Las plantas productoras de resina que dieron origen al ámbar de El Soplao, coníferas pertenecientes al menos a la familia extinta Cheirolepidiaceae y a otro grupo diferente por determinar, se desarrollarían cerca de la costa marina. Los incendios, como parte integrante de la dinámica del ecosistema, tuvieron un papel determinante en la producción y posterior acumulación de resina que dio lugar al yacimiento, provocando la secreción traumática de resina (en la que los insectos barrenadores de madera pudieron haber colaborado) y eliminando la cobertura vegetal del suelo del bosque. Ello facilitó que el agua de escorrentía erosionara el suelo y movilizara la resina depositada primariamente. Esta resina sufrió un corto transporte y se depositó de un modo paraauctóctono en las áreas de enterramiento definitivo, al no existir indicios de reelaboración ni retrabajamiento del ámbar. Estas áreas donde la resina quedó enterrada y fosilizó corresponden a ambientes de marjal costero o de estuario asociado a una llanura deltaica. Algunas masas de resina presentaron un largo periodo de exposición en las aguas salobres o marinas antes de enterrarse. Pese a hallarse en una fase de estudio inicial, la diversidad identificada de artrópodos en el ámbar de El Soplao es elevada, y en algunas casos ha brindado el registro más antiguo conocido de ciertos taxones. En un total de 549 inclusiones de artrópodos catalogadas, se han determinado 13 órdenes de hexápodos repartidos en unas 40 familias, cuatro órdenes de arácnidos y uno de crustáceos. La paleofauna de artrópodos de El Soplao muestra una serie de singularidades, tanto cuantitativas como cualitativas, en comparación con el resto de paleofaunas conocidas de los ámbares cretácicos. Estas singularidades paleofaunísticas tanto a nivel cuantitativo como cualitativo en el ámbar de El Soplao, aparte de los factores tafonómicos y teniendo además en cuenta los hallazgos anteriores en el ámbar de Peñacerrada I, podrían reflejar un cierto carácter endémico de los ecosistemas ibéricos durante el Albiense debido a la insularidad que la Placa Ibérica adquirió en el Jurásico Inferior. Existen tres especies de insectos comunes a los ámbares de El Soplao y Peñacerrada I (= Moraza). Ello sugiere una edad muy similar para ambos yacimientos de la Cuenca Vasco‐Cantábrica. Además, se han advertido ciertas similitudes entre las paleofaunas del ámbar de España (El Soplao y Peñacerrada I) (Albiense) y las del ámbar del Líbano (Barremiense‐ Aptiense), aunque se hallen espaciadas en el tiempo por varios millones de años. No obstante, es necesario aumentar las colecciones de los ámbares españoles para dar mayor fundamento a estas observaciones, así como revisar las dataciones del ámbar del Líbano. La elevada proporción de sininclusiones y muestras que brindan información paleoetológica o acerca de interacciones paleobióticas, tanto intraespecíficas como interespecíficas, dan al ámbar de El Soplao un elevado valor paleoecológico. Existen casos de posible enjambramiento, de cópula, de camuflage/defensa, de relación planta‐insecto, de parasitismo, de posible hematofagia y de posible relación depredador‐presa.The El Soplao amber outcrop (Rábago, Cantabria, Northern Spain), Early Cretaceous in age (about 110 Mya), has currently provided more than 500 arthropod inclusions. This PhD dissertation aimed to achieve in‐depth knowledge of the paleobiology associated with the arthropods from the El Soplao amber outcrop, allowing for a preliminary comparison with primary Cretaceous amber outcrops. This was possible only after a thorough systematic treatment of El Soplao arthropod inclusions that inferred their paleoautoecological, paleobiogeographical and paleoethological contexts to, ultimately, glean a holistic understanding of the paleoenvironmental setting. The resin‐producing plants responsible for El Soplao amber are conifers belonging to the extinct family Cheirolepidiaceae and, at least, another unknown conifer group. These trees would have lived close to the paleo‐coastline. Paleofires played a leading role in the production and accumulation of resin. The areas where the resin was buried and fossilized correspond to marshy or delta‐estuarine environments. Some resin pieces had a long exposure time to marine or brackish waters before being buried. Identified arthropod diversity is high, and the outcrop has provided the oldest known records for certain taxa. Thirteen hexapod orders (composed by about 40 families), four arachnid orders, and one crustacean order have been determined. The arthropod fauna from El Soplao is quantitatively and qualitatively singular when compared to faunas from other known Cretaceous ambers. Three insect species occur in both the El Soplao and Peñacerrada I (=Moraza) outcrops. The shared species between these two Basque‐Cantabrian Basin outcrops suggest they are similar in age. Moreover, similarities have been noted between the paleofauna from the Spanish amber outcrops (El Soplao and Peñacerrada I) (Albian) and the paleofauna from Lebanon (Barremian‐Aptian), despite these outcrops are separated in time by several million years. El Soplao amber is valuable in the abundance of syninclusions and samples that provide information about paleoethology and paleobiotic interactions. There are instances of possible swarming, mating, camouflage/defense, plant‐insect relationship, parasitism, possible hematophagy and possible predator‐prey relationship. An expanded English summary is provided at the end of the dissertation

The first unique-headed bug (Hemiptera, Enicocephalomorpha) from Cretaceous Iberian amber, and the Gondwanan connections of its palaeoentomological fauna

Enicocephalomorpha, also known as unique-headed bugs, are a seldom-collected infraorder of heteropteran insects whose evolutionary relationships have puzzled entomologists for more than a century. Unique-headed bugs are exceptionally rare in the fossil record, which hinders our understanding of the morphological transformations of the lineage across time and also affects the calibration of molecular clock estimates used to date the origins of the infraorder. Here, we report the discovery of Enicocephalinus ibericus sp. nov. from Iberian amber in the Ariño deposit in Spain, early Albian (Early Cretaceous) in age. The new species represents the second oldest fossil enicocephalomorphan to date, and the second record of this infraorder from European deposits. Remarkably, the closest relative of E. ibericus is the congeneric E. acragrimaldii Azar from Lebanese amber that is c. 20 myr older (Barremian), indicating a long-term persistence of the Enicocephalinus lineage across geological time. A review of the existing literature enabled us to record a total of 20 congeneric insect species that have been found in both Lebanese and Iberian ambers, suggesting the existence of previously underappreciated entomofaunal connections between southern Laurasia (the European archipelago) and northern Gondwana during the Cretaceous. We show that the palaeoentomological record holds remarkable potential for elucidating the faunistic exchanges and palaeobiogeographical patterns in the peri-Tethyan region during the Cretaceous

Early adaptations of true flies (Diptera) to moist and aquatic continental environments

Altres ajuts: CERCA Programme/Generalitat de CatalunyaInsect colonization of continental aquatic ecosystems and their immediate surroundings was paramount for the establishment of complex trophic nets and organic-matter recycling in those environments. True flies and other insects such as mayflies developed crucial ecological roles in early continental aquatic ecosystems, as early as the Triassic. However, the mode and tempo of these processes remain poorly known, partly due to a critical fossil record gap before the Middle Triassic. Here we study the dipterans from the early Middle Triassic Konservat-Lagerstätte of Pedra Alta (Aegean, early Anisian, Spain), which yields the oldest records of the order. Protoanisolarva juarezi gen. et sp. nov., based on an exceptionally preserved larva, shares key features with the extant nematoceran family Anisopodidae. Developing in inferred moist terrestrial environments contiguous with pools inhabited by aquatic organisms, it represents the only known Triassic dipteran larva with terrestrial affinities indicating that the amphipneustic respiratory system of insect larvae extends back to c. 247 Ma. Two nematoceran aquatic pupae are also described: one classified as Voltziapupa cf. cornuta, and the other as an indeterminate taxon. Finally, an egg cluster belonging to the ootaxon Clavapartus latus is likely to have been produced by chironomids. These eggs were included in a mucilaginous matrix, a probable adaptation against predation and/or changing conditions, including desiccation. These new findings provide key data on the early evolutionary history of the mega-diverse order Diptera, the ecology of their ancestral pre-adult forms, and the functioning of early Middle Triassic continental aquatic ecosystems

A defensive behavior and plant-insect interaction in Early Cretaceous amber - The case of the immature lacewing Hallucinochrysa diogenesi

Amber holds special paleobiological significance due to its ability to preserve direct evidence of biotic interactions and animal behaviors for millions of years. Here we review the finding of Hallucinochrysa diogenesi P erez-de la Fuente, Delcl os, Pe~nalver and Engel, 2012, a morphologically atypical larva related to modern green lacewings (Insecta: Neuroptera) that was described in Early Cretaceous amber from the El Soplao outcrop (northern Spain). The fossil larva is preserved with a dense cloud of fern trichomes that corresponds to the trash packet the insect gathered and carried on its back for camouflaging and shielding, similar to that which is done by its extant relatives. This finding supports the prominent role of wildfires in the paleoecosystem and provides direct evidence of both an ancient planteinsect interaction and an early acquisition of a defensive behavior in an insect lineage. Overall, the fossil of H. diogenesi showcases the potential that the amber record offers to reconstruct not only the morphology of fossil arthropods but, more remarkably, their lifestyles and ecological relationships

A new dustywing (Neuroptera: Coniopterygidae) from the Early Cretaceous amber of Spain.

A new Cretaceous dustywing, Soplaoconis ortegablancoi gen. et sp. nov. (Neuroptera: Coniopterygidae), is described from four specimens preserved in Early Cretaceous (Albian, ~105Ma) El Soplao amber (Cantabria, northern Spain). Two additional specimens are assigned to this new taxon. A crossvenational abnormality on an area of diagnostic significance from one of the holotype's forewings provides a reminder of the importance of not ruling out character plasticity or teratoses when evaluating palaeodiversity. A comment on the possible palaeoecological significance of the co-occurrence as syninclusions of plant trichomes with the holotype of S. ortegablancoi and seven of the eleven described Burmese amber dustywing species is provided

Absorption and translocation to the aerial part of magnetic carbon-coated nanoparticles through the root of different crop plants

The development of nanodevices for agriculture and plant research will allow several new applications, ranging from treatments with agrochemicals to delivery of nucleic acids for genetic transformation. But a long way for research is still in front of us until such nanodevices could be widely used. Their behaviour inside the plants is not yet well known and the putative toxic effects for both, the plants directly exposed and/or the animals and humans, if the nanodevices reach the food chain, remain uncertain. In this work we show that magnetic carbon-coated nanoparticles forming a biocompatible magnetic fluid (bioferrofluid) can easily penetrate through the root in four different crop plants (pea, sunflower, tomato and wheat). They reach the vascular cylinder, move using the transpiration stream in the xylem vessels and spread through the aerial part of the plants in less than 24 hours. Accumulation of nanoparticles was detected in wheat leaf trichomes, suggesting a way for excretion/detoxification. This kind of studies is of great interest in order to unveil the movement and accumulation of nanoparticles in plant tissues for assessing further applications in the field or laboratory

False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance.

During the mid-Cretaceous, angiosperms diversified from several nondiverse lineages to their current global domination [1], replacing earlier gymnosperm lineages [2].Several hypotheses explain this extensive radiation [3], one of which involves proliferation of insect pollinator associations in the transition from gymnosperm to angiosperm dominance. However, most evidence supports gymnosperm-insect pollinator associations, buttressed by direct evidence of pollen on insect bodies, currently established for four groups: Thysanoptera (thrips), Neuroptera (lacewings), Diptera (flies), and now Coleoptera (beetles). Each group represents a distinctive pollination mode linked to a unique mouthpart type and feeding guild [4-9]. Extensive indirect evidence, based on specialized head and mouthpart morphology, is present for one of these pollinator types, the long-proboscid pollination mode [10], representing minimally ten family-level lineages of Neuroptera, Mecoptera (scorpionflies), and Diptera [8, 10, 11]. A recurring feature uniting these pollinator modes is host associations with ginkgoalean, cycad, conifer, and bennettitalean gymnosperms. Pollinator lineages bearing these pollination modes were categorized into four evolutionary cohorts during the 35-million-year-long angiosperm radiation, each defined by its host-plant associations (gymnosperm or angiosperm) and evolutionary pattern (extinction, continuation, or origination) during this interval [12]. Here, we provide the first direct evidence for one cohort, exemplified by the beetle Darwinylus marcosi, family Oedemeridae (false blister beetles), that had an earlier gymnosperm (most likely cycad) host association, later transitioning onto angiosperms [13]. This association constitutes one of four patterns explaining the plateau of family-level plant lineages generally and pollinating insects specifically during the mid-Cretaceous angiosperm radiation[12]